Anthropogenic microfibres flux in an Antarctic coastal ecosystem: The tip of an iceberg?

This study describes the occurrence of anthropogenic microfibres (AMFs) found in sediment trap samples collected at 25 m water depth in an Antarctic fjord (Potter Cove, King George/25 de Mayo Island) from 2012 to 2015. During visual sorting of samples, AMFs were detected and described, and a subset was confirmed, via FTIR (Fourier transform infrared) spectroscopy, as semi-synthetic cellulosic and polyacrylonitrile polymers. Estimated flux of AMF varied from 115 to 152,750 microfibres m-2 throughout the studied period, with sizes ranging from 10 to 450 µm in length. Maximum AMFs fluxes occurred in summer months. Sediment traps allowed detecting temporal patterns of small (µm) AMFs, usually undersampled with nets or sieves, providing a new insight into microplastic pollution in Antarctica and its relation to environmental conditions.

Acute effect of a static- and dynamic-based stretching warm-up on standing long jump performance in primary schoolchildren.

The aim of the present study was to compare the acute effect of a static- vs dynamic-based stretching warm-up on standing long jump (SLJ) performance in primary schoolchildren. The sample was composed of 76 schoolchildren, 43 girls and 33 boys, aged 9-10 years old from three fourth-grade classes of Primary Education. The three groups were cluster-randomly assigned to the control (CG), static (SG) or dynamic (DG) groups. All the schoolchildren performed a standardized warm-up consisting of mobility exercises (five minutes), jogging (five minutes) and the SLJ test. Afterwards the CG schoolchildren received jump theory (eight minutes), the SG performed static stretching (eight minutes) and the DG performed dynamic-bounces stretching (eight minutes). Afterwards, all of them performed the SLJ test again. The results of the one-way ANOVA (F 2,73 = 34.184; p < 0.001; η2p = 0.484), followed by the pairwise comparisons with the Bonferroni adjustment, showed that the DG students (M Δ = 11.07, SE = 1.42) made a significant statistical improvement in their SJL levels compared with the CG (M Δ = -3.00, SE = 1.89; p < 0.001, d = 1.51) and SG students (M Δ = -1.85, SE = 0.67; p < 0.001, d = 1.38). However, statistically significant differences between the CG and SG students were not found (p > 0.05, d = 0.12). The dynamic-bouncing stretch as a final part of a warm-up improves explosive strength performance in primary schoolchildren, and seems to be a good option before carrying out explosive strength activities of the lower body.

Differing Effects of Vinegar on Pelagia noctiluca (Cnidaria: Scyphozoa) and Carybdea marsupialis (Cnidaria: Cubozoa) Stings-Implications for First Aid Protocols.

The jellyfish species that inhabit the Mediterranean coastal waters are not lethal, but their stings can cause severe pain and systemic effects that pose a health risk to humans. Despite the frequent occurrence of jellyfish stings, currently no consensus exists among the scientific community regarding the most appropriate first-aid protocol. Over the years, several different rinse solutions have been proposed. Vinegar, or acetic acid, is one of the most established of these solutions, with efficacy data published. We investigated the effect of vinegar and seawater on the nematocyst discharge process in two species representative of the Mediterranean region: Pelagia noctiluca (Scyphozoa) and Carybdea marsupialis (Cubozoa), by means of (1) direct observation of nematocyst discharge on light microscopy (tentacle solution assay) and (2) quantification of hemolytic area (tentacle skin blood agarose assay). In both species, nematocyst discharge was not stimulated by seawater, which was classified as a neutral solution. In P. noctiluca, vinegar produced nematocyst discharge per se, but inhibited nematocyst discharge from C. marsupialis. These results suggest that the use of vinegar cannot be universally recommended. Whereas in case of a cubozoan C. marsupialis sting, the inhibitory effect of vinegar makes it the ideal rinse solution, in case of a scyphozoan P. noctiluca sting, vinegar application may be counterproductive, worsening the pain and discomfort of the stung area.

Effects of environmental variables on the distribution of juvenile cubomedusae Carybdea marsupialis in the coastal Western Mediterranean.

Relationships between environmental factors and oscillations in jellyfish abundance, especially in the early life stages, could help to interpret past increases and also predict scenarios in a changing future. For the first time, we present cubozoan spatial and temporal distributions in the earliest stages and their relationships with different factors. Abundances of Carybdea marsupialis medusae showed high interannual variability from 2008 to 2014 along the Dénia coast (SE Spain, W Mediterranean). During 2015, samples were collected from 11 beaches along 17 km of coastline, 8 times from January to November in order to determine the effects of environmental factors on the distribution of juvenile C. marsupialis. Juveniles (≤ 15 mm diagonal bell width) were present from May to July, with more sampled near shore (0-15 m). Most of them occurred in June when their numbers were unequal among beaches (average 0.05 ind m-3, maximum 6.71 ind m-3). We tested distributions of juveniles over time and space versus temperature, salinity, nutrients (N, P and Si), chlorophyll-a (Chl-a), and zooplankton abundance. Temperature and cladocerans (zooplankton group) were significantly positively correlated with juvenile distribution, whereas Chl-a concentration was weakly negative. By contrast, in 2014, high productivity areas (Chl-a and zooplankton) overlapped the maximum adult abundance (5.2 ind m-3). The distribution of juveniles during 2015 did not spatially coincide with the areas where ripe adults were located the previous year, suggesting that juveniles drift with the currents upon release from the cubopolyps. Our results yield important insights into the complexity of cubozoan distributions.

Role of suspension feeders in antarctic pelagic-benthic coupling: Trophic ecology and potential carbon sinks under climate change.

Sea-ice and coastal glacier loss in the Western Antarctic Peninsula open new ice-free areas. They allowing primary production and providing new seabed for colonisation, both acting as a negative feedback of climate change. However, the injection of sediment-laden runoff from the melting of land-terminating glaciers may reduce this feedback. Changes in particulate matter will affect nutrition and excretion (faeces stoichiometry and properties) of suspension feeders, reshaping coastal carbon dynamics and pelagic-benthic coupling. Absorption efficiency and biodeposition of Euphausia superba and Cnemidocarpa verrucosa were quantified for different food treatments and varying sediment concentrations. Both species showed high overall absorption efficiency for free-sediment diets, but were negatively affected by sediment addition. High sediment conditions increased krill biodeposition, while it decreased in ascidians. Energy balance estimation indicated high carbon sink potential in ascidians, but it is modulated by food characteristics and negatively affected by sediment inputs in the water column.

Revision of the genus Carybdea (Cnidaria: Cubozoa: Carybdeidae): clarifying the identity of its type species Carybdea marsupialis.

While records of Carybdea marsupialis in the literature suggest a worldwide distribution of this species, the validity of some of these records has been questioned recently, as has the validity of some nominal Carybdea species. We inspected material of all known species of Carybdea from multiple locations (i.e. Spain, Algeria, Tunisia, Puerto Rico, California, Hawaii, Australia, South Africa, and Japan) using morphological and genetic tools to differentiate Carybdea species as well as understand their evolutionary relationships. We observed morphological differences between adult medusae of Mediterranean and Caribbean C. marsupialis; the most obvious differences were the structure of the phacellae, the structure of the pedalial canal knee bend, and the number and structure of the velarial canals. The characters of the adult Mediterranean specimens agree with the description provided by Claus (1878) for individuals of C. marsupialis from the Adriatic Sea (Italy); specimens from the Caribbean (Puerto Rico) agreed with the description of C. xaymacana by Conant (1897). Significant differences between both species were also observed in the newly released medusa stage. Further, we resolved a discord about the undefined polyp culture originating from Puerto Rico that was long considered Carybdea marsupialis but should be referred to as C. xaymacana. Although C. marsupialis is currently considered the only species of Cubozoa to occur in the Mediterranean, specimens collected in Algeria and Tunisia suggest that species of Alatinidae may also be present in the Mediterranean. Our investigations indicate that Carybdea spp. are more restricted in their geographical distribution than has been recognized historically. These findings confirm that Carybdea arborifera Maas, 1897 from Hawaii, Carybdea branchi, Gershwin Gibbons, 2009 from South Africa, Carybdea brevipedalia Kishinouye, 1891 from Japan, Carybdea confusa Straehler-Pohl, Matsumoto Acevedo, 2017 from California, Carybdea marsupialis Linnaeus, 1758 from the European Mediterranean Sea, Carybdea rastonii Haacke, 1886 from South Australia, and Carybdea xaymacana, Conant, 1897 from the Caribbean Sea are valid names representing distinct species, rather than synonyms. A taxonomic key for all valid species is provided, and a neotype for C. marsupialis is designated.

Seasonal variability of diet and trophic level of the gelatinous predator Pelagia noctiluca (Scyphozoa).

Jellyfish populations apparently have increased in some places around the world and human problems with them also have increased. However, effects of jellyfish outbreaks in the ecosystems remain poorly understood and little or no information is available on their dietary preferences - in relation to the seasonal shifts of prey abundance - and on the potential variability of their impact on marine food webs. The mauve stinger Pelagia noctiluca (Forsskål, 1775) is by far the most common outbreak-forming scyphozoan jellyfish in the Western Mediterranean. By use of a combination of stomach contents, stable isotope (SI) and fatty acid (FA) analyses, we tested the hypothesis that changes in the seasonal dietary sources of P. noctiluca parallel changes in the FA and SI composition. Stomach content and biomarker analyses suggested that P. noctiluca is not a selective predator, cyclically shifting between carnivory and omnivory depending on the seasonality of accessible prey. The combination of SI and FA analyses highlighted the importance of microzooplankton as prey. Specific FA biomarkers showed that the diet of P. noctiluca changed seasonally depending on the availability of living plankton or suspended detritus. This study also revealed significant biochemical differences between jellyfish somatic and gonadal tissues, with total fatty acid concentration in the gonads up to ten times higher than in the somatic tissues.

Environmental factors influencing the spatio-temporal distribution of Carybdea marsupialis (Lineo, 1978, Cubozoa) in South-Western Mediterranean coasts.

Jellyfish blooms cause important ecological and socio-economic problems. Among jellyfish, cubozoans are infamous for their painful, sometimes deadly, stings and are a major public concern in tropical to subtropical areas; however, there is little information about the possible causes of their outbreaks. After a bloom of the cubomedusa Carybdea marsupialis (Carybdeidae) along the coast of Denia (SW Mediterranean, Spain) in 2008 with negative consequences for local tourism, the necessity to understand the ecological restrictions on medusae abundance was evident. Here we use different models (GAM and zero-inflated models) to understand the environmental and human related factors influencing the abundance and distribution of C. marsupialis along the coast of Denia. Selected variables differed among medusae size classes, showing different environmental restriction associated to the developmental stages of the species. Variables implicated with dispersion (e.g. wind and current) affected mostly small and medium size classes. Sea surface temperature, salinity and proxies of primary production (chl a, phosphates, nitrates) were related to the abundances of small and large size classes, highlighting the roles of springtime salinity changes and increased primary production that may promote and maintain high densities of this species. The increased primary (and secondary) production due to anthropogenic impact is implicated as the factor enabling high numbers of C. marsupialis to thrive. Recommendations for monitoring blooms of this species along the study area and applicable to Mediterranean Sea include focus effort in coastal waters where productivity have been enriched by anthropogenic activities.

Glacial melting: an overlooked threat to Antarctic krill.

Strandings of marine animals are relatively common in marine systems. However, the underlying mechanisms are poorly understood. We observed mass strandings of krill in Antarctica that appeared to be linked to the presence of glacial meltwater. Climate-induced glacial meltwater leads to an increased occurrence of suspended particles in the sea, which is known to affect the physiology of aquatic organisms. Here, we study the effect of suspended inorganic particles on krill in relation to krill mortality events observed in Potter Cove, Antarctica, between 2003 and 2012. The experimental results showed that large quantities of lithogenic particles affected krill feeding, absorption capacity and performance after only 24 h of exposure. Negative effects were related to both the threshold concentrations and the size of the suspended particles. Analysis of the stomach contents of stranded krill showed large quantities of large particles ( > 10(6 )µm(3)), which were most likely mobilized by glacial meltwater. Ongoing climate-induced glacial melting may impact the coastal ecosystems of Antarctica that rely on krill.

Concurrent environmental stressors and jellyfish stings impair caged European sea bass (Dicentrarchus labrax) physiological performances.

The increasing frequency of jellyfish outbreaks in coastal areas has led to multiple ecological and socio-economic issues, including mass mortalities of farmed fish. We investigated the sensitivity of the European sea bass (Dicentrarchus labrax), a widely cultured fish in the Mediterranean Sea, to the combined stressors of temperature, hypoxia and stings from the jellyfish Pelagia noctiluca, through measurement of oxygen consumption rates (MO2), critical oxygen levels (PO2crit), and histological analysis of tissue damage. Higher levels of MO2, PO2crit and gill damage in treated fish demonstrated that the synergy of environmental and biotic stressors dramatically impair farmed fish metabolic performances and increase their health vulnerability. As a corollary, in the current scenario of ocean warming, these findings suggest that the combined effects of recurrent hypoxic events and jellyfish blooms in coastal areas might also threaten wild fish populations.

Jellyfish Stings Trigger Gill Disorders and Increased Mortality in Farmed Sparus aurata (Linnaeus, 1758) in the Mediterranean Sea.

Jellyfish are of particular concern for marine finfish aquaculture. In recent years repeated mass mortality episodes of farmed fish were caused by blooms of gelatinous cnidarian stingers, as a consequence of a wide range of hemolytic, cytotoxic, and neurotoxic properties of associated cnidocytes venoms. The mauve stinger jellyfish Pelagia noctiluca (Scyphozoa) has been identified as direct causative agent for several documented fish mortality events both in Northern Europe and the Mediterranean Sea aquaculture farms. We investigated the effects of P. noctiluca envenomations on the gilthead sea bream Sparus aurata by in vivo laboratory assays. Fish were incubated for 8 hours with jellyfish at 3 different densities in 300 l experimental tanks. Gill disorders were assessed by histological analyses and histopathological scoring of samples collected at time intervals from 3 hours to 4 weeks after initial exposure. Fish gills showed different extent and severity of gill lesions according to jellyfish density and incubation time, and long after the removal of jellyfish from tanks. Jellyfish envenomation elicits local and systemic inflammation reactions, histopathology and gill cell toxicity, with severe impacts on fish health. Altogether, these results shows P. noctiluca swarms may represent a high risk for Mediterranean finfish aquaculture farms, generating significant gill damage after only a few hours of contact with farmed S. aurata. Due to the growth of the aquaculture sector and the increased frequency of jellyfish blooms in the coastal waters, negative interactions between stinging jellyfish and farmed fish are likely to increase with the potential for significant economic losses.

Effects of the harmful dinoflagellate Ostreopsis cf. ovata on different life cycle stages of the common moon jellyfish Aurelia sp.

The frequency and geographic extension of microalgae and gelatinous zooplankton blooms seem to have been increasing worldwide over recent decades. In particular, the harmful dinoflagellate Ostreopsis cf. ovata and the Schyphozoan jellyfish Aurelia sp. are two of the most frequent and long lasting species forming blooms in the Mediterranean Sea. A kind of interaction among any of their life cycle stages (i.e. planula-polyp-ephyrae vs Ostreopsis cells) can likely occur, although in this area there are no data available on the co-occurrence of these species. The aim of this study was to investigate, for the first time, the potential noxious effect of O. cf. ovata on different life stages of Aurelia sp. (polyps and ephyrae), testing several concentrations of whole algal culture. Rsults of toxicity bioassay highlighted that ephyrae, but not polyps, are affected by this harmful dinoflagellate and comparisons among other model organisms show that Aurelia sp. ephyrae are the most sensitive model organism tested so far (EC50-24h=10.5cells/mL). These findings suggest an interesting scenario on the interaction of these two bloom forming species in the natural marine environment.

Deterministic Factors Overwhelm Stochastic Environmental Fluctuations as Drivers of Jellyfish Outbreaks.

Jellyfish outbreaks are increasingly viewed as a deterministic response to escalating levels of environmental degradation and climate extremes. However, a comprehensive understanding of the influence of deterministic drivers and stochastic environmental variations favouring population renewal processes has remained elusive. This study quantifies the deterministic and stochastic components of environmental change that lead to outbreaks of the jellyfish Pelagia noctiluca in the Mediterranen Sea. Using data of jellyfish abundance collected at 241 sites along the Catalan coast from 2007 to 2010 we: (1) tested hypotheses about the influence of time-varying and spatial predictors of jellyfish outbreaks; (2) evaluated the relative importance of stochastic vs. deterministic forcing of outbreaks through the environmental bootstrap method; and (3) quantified return times of extreme events. Outbreaks were common in May and June and less likely in other summer months, which resulted in a negative relationship between outbreaks and SST. Cross- and along-shore advection by geostrophic flow were important concentrating forces of jellyfish, but most outbreaks occurred in the proximity of two canyons in the northern part of the study area. This result supported the recent hypothesis that canyons can funnel P. noctiluca blooms towards shore during upwelling. This can be a general, yet unappreciated mechanism leading to outbreaks of holoplanktonic jellyfish species. The environmental bootstrap indicated that stochastic environmental fluctuations have negligible effects on return times of outbreaks. Our analysis emphasized the importance of deterministic processes leading to jellyfish outbreaks compared to the stochastic component of environmental variation. A better understanding of how environmental drivers affect demographic and population processes in jellyfish species will increase the ability to anticipate jellyfish outbreaks in the future.

Use of an Inverse Method for Time Series to Estimate the Dynamics of and Management Strategies for the Box Jellyfish Carybdea marsupialis.

Frequently, population ecology of marine organisms uses a descriptive approach in which their sizes and densities are plotted over time. This approach has limited usefulness for design strategies in management or modelling different scenarios. Population projection matrix models are among the most widely used tools in ecology. Unfortunately, for the majority of pelagic marine organisms, it is difficult to mark individuals and follow them over time to determine their vital rates and built a population projection matrix model. Nevertheless, it is possible to get time-series data to calculate size structure and densities of each size, in order to determine the matrix parameters. This approach is known as a "demographic inverse problem" and it is based on quadratic programming methods, but it has rarely been used on aquatic organisms. We used unpublished field data of a population of cubomedusae Carybdea marsupialis to construct a population projection matrix model and compare two different management strategies to lower population to values before year 2008 when there was no significant interaction with bathers. Those strategies were by direct removal of medusae and by reducing prey. Our results showed that removal of jellyfish from all size classes was more effective than removing only juveniles or adults. When reducing prey, the highest efficiency to lower the C. marsupialis population occurred when prey depletion affected prey of all medusae sizes. Our model fit well with the field data and may serve to design an efficient management strategy or build hypothetical scenarios such as removal of individuals or reducing prey. TThis This sdfsdshis method is applicable to other marine or terrestrial species, for which density and population structure over time are available.

Carybdea marsupialis (Cubozoa) in the Mediterranean Sea: the first case of a sting causing cutaneous and systemic manifestations.

A woman stung by the box jellyfish Carybdea marsupialis (Cnidaria, Cubozoa) at a Spanish Mediterranean beach showed systemic manifestations over several months [pain far from the inoculation point, arthralgia, paresthesia, hyperesthesia, increase in eosinophils and immunoglobulin E (IgE)] in addition to the skin condition.

Jellyfish as prey: frequency of predation and selective foraging of Boops boops (Vertebrata, Actinopterygii) on the mauve stinger Pelagia noctiluca (Cnidaria, Scyphozoa).

In recent years, jellyfish blooms have attracted considerable scientific interest for their potential impacts on human activities and ecosystem functioning, with much attention paid to jellyfish as predators and to gelatinous biomass as a carbon sink. Other than qualitative data and observations, few studies have quantified direct predation of fish on jellyfish to clarify whether they may represent a seasonally abundant food source. Here we estimate predation frequency by the commercially valuable Mediterranean bogue, Boops boops on the mauve stinger jellyfish, Pelagia noctiluca, in the Strait of Messina (NE Sicily). A total of 1054 jellyfish were sampled throughout one year to quantify predation by B. boops from bite marks on partially eaten jellyfish and energy density of the jellyfish. Predation by B. boops in summer was almost twice that in winter, and they selectively fed according to medusa gender and body part. Calorimetric analysis and biochemical composition showed that female jellyfish gonads had significantly higher energy content than male gonads due to more lipids and that gonads had six-fold higher energy content than the somatic tissues due to higher lipid and protein concentrations. Energetically, jellyfish gonads represent a highly rewarding food source, largely available to B. boops throughout spring and summer. During the remainder of the year, when gonads were not very evident, fish predation switched towards less-selective foraging on the somatic gelatinous biomass. P. noctiluca, the most abundant jellyfish species in the Mediterranean Sea and a key planktonic predator, may represent not only a nuisance for human leisure activities and a source of mortality for fish eggs and larvae, but also an important resource for fish species of commercial value, such as B. boops.

Impact of high pressure treatment and intramuscular fat content on colour changes and protein and lipid oxidation in sliced and vacuum-packaged Iberian dry-cured ham.

The effect of high hydrostatic pressure (HHP) (600MPa) and intramuscular fat content (IMF) on colour parameters and oxidative stability of lipids and proteins in sliced vacuum-packaged Iberian dry-cured ham during refrigerated storage (120 days at 2°C) was investigated. Several studies have investigated the influence of HHP on lipid oxidation of meat products. However, its effects on protein carbonylation, as also the influence of IMF content on this carbonylation are poorly understood. HHP treatment had a significant effect on lean lightness after 0 and 120 days of storage while IMF content increased lightness and yellowness over time. Regarding oxidative stability, the effect of HHP treatment depended on IMF content samples with a high IMF having greater lipid instability while samples with a low IMF underwent more protein carbonylation.

The genetic background affects composition, oxidative stability and quality traits of Iberian dry-cured hams: purebred Iberian versus reciprocal Iberian × Duroc crossbred pigs.

This study examined the physico-chemical characteristics, oxidative stability and sensory properties of Iberian cry-cured hams as affected by the genetic background of the pigs: purebred Iberian (PBI) pigs vs reciprocal cross-bred Iberian × Duroc pigs (IB × D pigs: Iberian dams × Duroc sires; D × IB pigs: Duroc dams × Iberian sires). Samples from PBI pigs contained significantly higher amounts of IMF, monounsaturated fatty acids, heme pigments and iron than those from crossbred pigs. The extent of lipid and protein oxidation was significantly larger in dry-cured hams of crossbred pigs than in those from PBI pigs. Dry-cured hams from PBI pigs were defined by positive sensory properties (i.e. redness, brightness and juiciness) while hams from crossbred pigs were ascribed to negative ones (i.e. hardness, bitterness and sourness). Hams from PBI pigs displayed a superior quality than those from crossbred pigs. The position of the dam or the sire in reciprocal Iberian × Duroc crosses had no effect on the quality of Iberian hams.

Impact of lipid content and composition on lipid oxidation and protein carbonylation in experimental fermented sausages.

This study aims to investigate the effect of lipid content (∼4%, ∼10% and ∼15%) and composition (different lipid sources; animal fat and sunflower oil) on the oxidative stability of proteins and lipids in experimental fermented sausages. Increasing the lipid content of sausages enhanced the susceptibility of lipids to oxidation whereas the effect on the formation of specific carbonyls from protein oxidation was not so evident. Sausages manufactured with different lipid sources affected the susceptibility of lipids and proteins to oxidation as a likely result of the modifications in the fatty acid profile, as well as to the presence of antioxidant compounds. While the fatty acid profile had a major effect on the occurrence and extent of lipid oxidation, the presence of compounds with potential antioxidant activity may be more influential on the extent of protein carbonylation.

Invasion pathway of the Ctenophore Mnemiopsis leidyi in the Mediterranean Sea.

Gelatinous zooplankton outbreaks have increased globally owing to a number of human-mediated factors, including food web alterations and species introductions. The invasive ctenophore Mnemiopsis leidyi entered the Black Sea in the early 1980s. The invasion was followed by the Azov, Caspian, Baltic and North Seas, and, most recently, the Mediterranean Sea. Previous studies identified two distinct invasion pathways of M. leidyi from its native range in the western Atlantic Ocean to Eurasia. However, the source of newly established populations in the Mediterranean Sea remains unclear. Here we build upon our previous study and investigate sequence variation in both mitochondrial (Cytochrome c Oxidase subunit I) and nuclear (Internal Transcribed Spacer) markers in M. leidyi, encompassing five native and 11 introduced populations, including four from the Mediterranean Sea. Extant genetic diversity in Mediterranean populations (n = 8, N a = 10) preclude the occurrence of a severe genetic bottleneck or founder effects in the initial colonizing population. Our mitochondrial and nuclear marker surveys revealed two possible pathways of introduction into Mediterranean Sea. In total, 17 haplotypes and 18 alleles were recovered from all surveyed populations. Haplotype and allelic diversity of Mediterranean populations were comparable to populations from which they were likely drawn. The distribution of genetic diversity and pattern of genetic differentiation suggest initial colonization of the Mediterranean from the Black-Azov Seas (pairwise F ST = 0.001-0.028). However, some haplotypes and alleles from the Mediterranean Sea were not detected from the well-sampled Black Sea, although they were found in Gulf of Mexico populations that were also genetically similar to those in the Mediterranean Sea (pairwise F ST = 0.010-0.032), raising the possibility of multiple invasion sources. Multiple introductions from a combination of Black Sea and native region sources could be facilitated by intense local and transcontinental shipping activity, respectively.